Natural wounds are produced in a variety of ways during the life of the plant, and, generally speaking, are easily healed over by the normal process if the area destroyed is not too large, and the parts remaining uninjured are sufficiently provided with foliage, or with supplies of food-materials stored up in the roots, rhizomes, medullary rays, etc., to feed a vigorous callus.
The nature of such wounds and the mode of healing are explained by what we know of artificial wounds, and it only remains to point out that the princ.i.p.al danger of ordinary wounds is not so much the direct traumatic action, because the simpler organisation of the plant does not involve matters connected with shock, loss of blood, etc., as in animals; the danger consists, rather, in their affording access to other injurious agents, especially fungi, and the treatment of wounds frequently resolves itself into cutting or pruning in order to get clean surfaces which can heal readily.
Wounds on leaves imply loss of foliar surface--_i.e._ of chlorophyll action--and the remarks on page 193 apply.
_Burrows_ may be taken as comprising all kinds of tunnel-like excavations in the various organs of plants, including those cases where insects burrow into hollow stems of gra.s.ses, etc., as indicated by the perforations they make in the outer tissues.
_Bark-boring_ is done by many species of beetles, especially _Scolytidae_, which excavate characteristically formed branching pa.s.sages tangentially in the inner bark of Conifers and other trees.
Some of them also bore down to the surface of the sap wood (_e.g._ _Tomicus bidentatus_) or even burrow right into the latter (_e.g._ _T.
lineatum_). It commonly happens that the external apertures show up clearly, owing to the brown dust and excrement, sometimes accompanied by turpentine, which exude from them. Many of these Bark beetles only attack trees which are already injured by fire, lightning, etc.; possibly they cannot bore through a cortex which swamps them with sap, as a vigorous one might do.
_Wood-boring_ is also done by many of the bark-beetles as well as by Longicorns, _e.g._ _Saperda_ in Poplars and Willows, the young shoots of which often show characteristic swellings with lateral holes indicating the points of exit. From the external apertures comminuted wood, like saw-dust, is frequently ejected in quant.i.ty and betrays the presence of the insects. Certain wood-wasps (_Sirex_) and the larvae of moths (_Cossus_) also make large perforations in the wood of Willows and other trees, often destroying it completely. In the case of these larger borers, whose tunnels may be as broad as the little finger, the foul smell as well as abundant "saw-dust" betray the evil.
Excavations in wood are by no means caused only by insects: several of the larger Hymenomycetes--_Stereum_, _Thelephora_, _Polyporus_, etc.--tunnel the timber in characteristic ways and often after a fashion very suggestive of insects. They usually obtain access through fractures.
_Tunnels_ in leaves are invariably due to the activity of miners belonging to the smaller moths and beetles--_e.g._ _Tinea_, _Orchestes_, etc.--the larvae of which eat out the mesophyll but leave the covering epidermis or cuticle untouched, and since the insect bores forwards only, in an irregular track, and leaves its excrement in the winding pa.s.sage, the effect is very characteristic.
Whitish leaf tunnels in Peas are excavated by _Phytomyza_.
Characteristic foxy-red tunnels are mined in the leaves of Apples by _Lyonetia_, _Coleophora_, etc.
_Falling of fruit_, of Apples, Plums, Apricots, etc., before they are ripe, is frequently due to insects, of which the various species of _Grapholitha_ or _Carpocapsa_ are conspicuous: the fallen fruits show a small hole leading by a labyrinth of pa.s.sages to the "core" or "stone,"
and in which the grub and its excrement are visible. The cutting off of the vascular bundles and disturbance of the water supply only partly explain the premature fall.
_Pith-flecks_ are minute brown specks or patches found in the wood-layers of many trees, and consist of dead parenchymatous thick-walled cells, reminding one of the structure of pith. They are explained as due to the borings of minute insects, _Diptera_ or Beetles, the larvae of which pierce the cortex and phloem and bore their way into the cambium. The latter then occludes the tunnels by filling them up with cells, and continuing its wood-forming activity gradually buries them deeper and deeper in the wood. Such pith-flecks are common in Willow, Birch, Alder, _Sorbus_, etc. It is possible that they may be due to other causes also in other trees.
_Erosions_ or _irregular wounds_ on leaves are caused by large numbers of grubs and caterpillars and other insects, such as earwigs, as well as slugs, snails, and other animals; but it must by no means be a.s.sumed that all marginal leaf wounds, for instance, are caused by animals, since many fungi which rot the tissues, as explained below (p. 208), also cause such erosions, the putrescent parts falling out--_e.g._ the Potato disease.
_Skeleton leaves_ frequently result from the ravages of caterpillars, which leave the coa.r.s.er ribs and veins untouched, but much finer skeletons with the minute veins almost intact may be found on plants infested with certain insects--_e.g._ _Selandria_ on Cherries.
Skeletonised patches on Cherry leaves, often pink or brown-pink, are eaten out by this grub.
_Shot-holes_ are perforations in leaves presenting the appearance, from their more or less rounded shape, of gunshot wounds. They may be due to insects which bore through the young leaves while still folded in the bud--_e.g._ Willow Beetle--or which gnaw out the tissue--_e.g._ the Beech Miner. Similar but usually more torn and irregular holes are eaten out by many caterpillars--_e.g._ the Cabbage Moth.
Shot-holes on Peas may be the work of Thrips.
Leaf perforations are commonly caused by severe hail-storms, the hail-stones beating right through the thin mesophyll. Certain chemicals used for spraying have also been known to cause shot-holes by killing the tissue beneath the standing drops.
There is, however, a cla.s.s of shot-holes in thin leaves which are due to the action of minute fungi, the mycelium of which so rots the tissues in a more or less circular area round the point of infection, that, in wet weather, the decomposing ma.s.s falls out and leaves a round hole--_e.g._ certain Chytridiaceae, Peronosporeae, _Gloeosporium_, _Exoascus_, etc.
If dry weather supervenes these holes frequently dry at the edges, and the leaves appear as if eaten out.
Shot-holes in Cherry, Walnut, Tobacco, and Plum leaves are due to _Phyllosticta_, in Cherry leaves also to _Clasterosporium_, and in Potato leaves to _Haltica_.
_Frost-cracks._--The trunks of trees exposed to the north-east, and occasionally with other aspects, are apt to show longitudinal ridges which realise on a larger scale the features of healed wounds scored with a knife. These wounds are due to the outer layers of wood losing water from their cell-walls as it congeals to ice in their lumina, more rapidly than do the warmer internal parts of the trunk; as this drying of the wood causes its shrinkage, especially in the tangential direction, the effect of a sudden frost and north-east wind is to rend the wood, which splits longitudinally with a loud report, as may often be heard in severe winters. Since the cortex and bark are ruptured at the same time the total effect resembles that of a deep knife-cut, and the same healing processes result on a larger scale when the wood swells and closes up the wound again in spring. But this recently-closed lesion is evidently a plane of weakness, and if a similarly severe winter follows the wound reopens and again heals, and so on, until after a succession of years a prominent _Frost-ridge_ results, which may finally heal completely if milder winters ensue or the tree be eventually protected.
_Strangulations._--We are now in a position to understand the so-called strangulations which result when woody climbers, telegraph wires, etc., kill or injure trees by tightly winding round them. If strong wire is twisted horizontally round a stem, the growth in thickness of the latter causes the trapping of the cortex and cambium, etc., between the wire and the wood, and a ringing process is set up in consequence of the death of the compressed tissues. A callus then forms above the wound, as in the case of true ringing by means of a cut, and eventually bulges over the upper side of the wire: in the course of years this overgrowth may completely cover in the wire, and, pressing on to the lower lip of the wound, may at length fuse with the cambium below. Hereafter the thickening rings of wood are continuous over the buried wire. The process is obstructed by all the impediments referred to in dealing with ringing, and of course the stem thickens more above than below the wire.
If the sapwood is thin, and the bark is so thick as to put great obstacles in the way of the junction of the upper and lower cambiums, death may result--the tree is permanently ringed. (See p. 201.)
_Spiral grooves_ are frequently met with where Wood-bine or other woody climbers have twined round a young stem or branch, the upper lip of the groove always protruding more than the lower. If a kink or a crossing of two plants or branches of the twiner results in a complete horizontal ring, the results are as in the above cases of ringing and strangulation. Naturally grooved walking sticks are often seen.
_Buried letters, etc._--These processes of healing by occlusion enable us to understand how letters of the alphabet, cut into the wood of trees, come to be buried deep in the timber as successive annual rings cover them in more and more. Chains, nails, rope, etc., have frequently been found thus buried in wood.
NOTES TO CHAPTER XXII.
In addition to the notes to the last chapter, the reader may be referred to Fisher in Vol. IV. of Schlich"s _Manual of Forestry_, Chap. VI., for an account of Hess" excellent work on Boring Beetles, etc.
The authority on Wood-fungi is Hartig, see especially his _Zersetzungs-erscheinungen des Holzes_, the princ.i.p.al results of which are condensed in his _Diseases of Trees_ already referred to. As regards "Pith-flecks," the reader should consult Frank, _Krankh. der Pflanzen_, B. I., p. 212: the subject needs further investigation.
CHAPTER XXIII.
EXCRESCENCES.
_Herbaceous excrescences, or galls--Erineum--Intumescences-- Corky warts, etc.--Pustules--Frost-blisters--Galls and Cecidia --Root nodules._
_Excrescences_, or out-growths of more or less abnormal character from the general surface of diseased organs, are very common symptoms, and widely recognised. They are due to hypertrophy of the tissues while the cells are young and capable of growth, and may be induced by a variety of causes, among which the stimulus of insect-punctures and of the presence of insect eggs are best known; but that of fungi, though less widely recognised, plays an equally important part, and, as we shall see, galls and other excrescences may be due to widely different agents.
_Galls_ or _Cecidia_ are protuberances of the most varied shapes, colours, and sizes found on herbaceous parts attacked by insects, fungi, etc. In the simplest cases the insects only pierce and suck the young cellular tissue--_e.g._ _Phytoptus_, Aphides, etc.--but in others the stimulus to hypertrophy starts by the puncture of the embryonic tissue of a leaf, root, etc., by the ovipositor of the female insect, which then lays an egg--_e.g._ _Cynips_, _Cecidomyia_, etc.--the presence of which appears to intensify the irritating action, or such only occurs when the young larva escapes.
Our knowledge of the primary cause of gall-formation amounts to very little. Generally speaking, only embryonic or very young cellular tissue reacts, and galls on adult leaves and branches have usually been initiated long before. The same gall-insect may induce totally different galls on different plants, or even on different parts of the same plant, and different insects call forth different galls on any one plant. These facts point clearly to the co-operation of both plant and insect in the gall-formation, and the best hypothesis yet to hand is to the effect that a gall is a hypertrophy of cells, the normal nutrition, growth, and division of which have been disturbed owing to the action of some poison or other irritant derived from the insect, or fungus, or other organism.
Attempts have been made to reproduce galls by injecting the juices of similar galls into the tissue, but as yet without success, and this may point to the co-operation of mechanical irritation during the hypertrophy in normal gall-formation.
Galls, in the broad sense, are not always preceded by a wound, however.
Insects on the outside of young tissues may cause such irritations that the parts in contact with the animal are arrested in their growth, while those further away grow more rapidly--_e.g._ where Mites, etc., cause puckers and leaf-rolling. In true galls the hypertrophy may consist merely in the enlargement of cells already present, and no new cell-divisions and, still less, changes in the nature of the tissues result--_e.g._ some pocket galls on _Viburnum_, _Pyrus_, etc., and the hairy outgrowths of the epidermis known as _Erineum_. In other cases there is not only hypertrophy of existing cells, but new cell-divisions are inst.i.tuted: these cell-divisions may be confined to the direction perpendicular to the epidermis, and the tissues grow only in the direction of the surface, producing puckerings--_e.g._ the Aphis galls on _Ribes_, Phytoptus galls of _Salvia_, leaf galls on _Tilia_, _Acer_, _Alnus_, etc., and the curious galls on Plums due to _Cecidomyia Pruni_, and which must not be confounded with the "pocket plums" and similar galls due to Exoasci.
In a third series of cases, cell-divisions occur parallel to the surface of the leaf, and galls are formed which grow in thickness, and develop the most extraordinary and complicated new tissues--proteid-cells surrounding the egg or larva deposited inside, followed by a protective layer of sclerenchyma encasing this food layer, and around this again softer tissues which may a.s.sume the structures and functions of respiratory tissues, water-storing tissues, starch reservoirs, a.s.similatory, or protective tissues of various kinds, and over all may be a well-marked epidermis, with stomata, or cork with lenticels.
The chief seat of these hypertrophies and--what is more remarkable--development of new tissue elements not found elsewhere in the leaves, or even in the species, is the mesophyll, and various speculations and hypothesis have been founded on these curious phenomena.
_Erineum._--The simplest excrescences on plants are certain hair-like developments of epidermal cells due to the irritation of species of _Phytoptus_, and similar insects which rise in cl.u.s.ters on the surfaces of leaves and by their colours, consistence, arrangement in patches, spots, etc., so simulate fungi that Persoon was deceived by them and gave them the genus name _Erineum_. They occur on most of our trees, _e.g._ Poplar, Lime, Oak, and are very common in the Tropics. Usually pale or even white at first, they turn brown as the hair-like outgrowths die and lose their sap, but since the latter may be bright coloured--yellow, red, purple,--the patches are sometimes very conspicuous objects on smooth leaves.
In many cases these hairs exactly resemble in shape and other characters the abnormal root-hairs found on roots exposed to the effects of poisonous reagents, or of unsuitable food-materials, or the rhizoids developed from wounded Algae, etc.
_Intumescences_ are similar trichomatous outgrowths not a.s.sociated with insects or fungi, and due to some disturbance of the balance between transpiratory and a.s.similatory functions of their leaves, as indicated by the less localised occurrence and by their non-appearance when the plant is under favourable cultural conditions. Structures not unlike these have been artificially induced by exposure to particular lights, and also by painting spots with dilute corrosive sublimate, indicating that poisons may impel the epidermis cells to grow out abnormally.
_Corky warts._--Several forms of disease are known in which the pathological condition is expressed by the formation of cork in unwonted places and quant.i.ties. The _Scab_ or _Scurf_ of Potatoes is a case in point. The tissue of the lenticels absorbs water and the outermost cells are cut off by cork and die: the cells below them burst the dead bark-like ma.s.ses thus formed, and again cork is formed and cuts off the outer ma.s.ses, and the rough cork warts--_Scab_ or _Scurf_--are the result.
The causes predisposing to scab have been variously a.s.signed to dampness, want of lime, action of bacteria and fungi--_e.g._ _Sorosporium_, _Oospora_, _Spongospora_,--the latter making their way into the ruptured tissue of the lenticels and irritating the cells to further growth.
It seems probable that several different kinds of scab exist in Potatoes, as well as in roots--_e.g._ Beets, and the whole subject needs further investigation. The scab-like rough scaly bark of Pear trees in dry districts may also be mentioned here.
_Cork-wings_ are well known on the young branches of Elms, Maples, etc., some varieties of which have received specific names on this account.
_Corky excrescences_ on leaves occur occasionally in the Gooseberry, Holly and other plants, for which no cause has been discovered.